Die Vererbung der Kernlosigkeit auf die Nachkommen der Kreuzungskombination „kernhaltig x kernlos" bei Vitis vinifera L.In 5jährigen Versuchen wurden kernhaltige (Mutter) und kernlose (Vater) Vitis vinifera-Reben gekreuzt und die Verteilung des Merkmals „Kernlosigkeit" bei den Nachkommen analysiert. Die Aufspaltung in kernhaltige und kernlose Sämlinge erfolgte nahezu im Verhältnis 3 : 1. Der x2-Wert für die einzelnen Jahre (1983, 1984, 1985, 1986, 1988) betrug 0,03; 0,11; 1,17; 1,97 und 3,01. Die Homogenität für x2 lag nur 1986 innerhalb der tolerierbaren Grenzen (0,3-0,5). Die meisten Populationen hatten in allen Jahren x2-Werte, die dem 3: 1-Verhältnis entsprachen. Zwei Independenz-Tests für aufeinanderfolgende Jahre ergaben keine Interaktionen. Auf der Grundlage dieser Ergebnisse wird postuliert, daß die Vererbung der Kernlosigkeit durch zwei komplementäre rezessive Gene bestimmt wird. Nur bei 30,7 % der als ,,kernlos" klassifizierten Kreuzungsnachkommen konnten die Kernreste organoleptisch nicht wahrgenommen werden

Baron, Y.

Sahar, N.

Spiegel-Roy, P.

English

JKI Open Journal Systems (Julius Kühn-Institut)

Vitis 29, 79-83 ( 1990) Department of Fruit Breeding and Genetics, Volcani Center, ARO, Bet Dagan, Israel Inheritance of seedlessness in seeded x seedless progeny of Vitis vinifera L. by P. SPIEGEL-ROY, Y. BARON and N. SAHAR Die Vererbung der Kernlosigkeit auf die Nachkommen der Kreuzungskombination „kernhaltig x kernlos" bei Vitis vinifera L. Zusammenfassung: In 5jährigen Versuchen wurden kernlialtige (Mutter) und kern-lose (Vater) Vitis vinifera-Reben gekreuzt und die Verteilung des Merkmals „Kernlosigkeit" bei den Nachkommen analysiert. Die Aufspaltung in kernhaltige und kernlose Sämlinge erfolgte nahezu im Verhältnis 3 : 1. Der x,2-Wert für die einzelnen Jahre (1983, 1984, 1985, 1986, 1988) betrug 0,03; 0,11; 1,17; 1,97 und 3,01. Die Homogenität für x,2 lag nur 1986 innerhalb der tolerierbaren Grenzen (0,3-0,5). Die meisten Populationen hatten in allen Jahren x,2-Werte, die dem 3: 1-Verhältnis ent-sprachen. Zwei Independenz-Tests für aufeinanderfolgende Jahre ergaben keine Interaktionen. Auf der Grundlage dieser Ergebnisse wird postuliert, daß die Vererbung der Kernlosigkeit durch zwei komplementäre rezessive Gene bestimmt wird. Nur bei 30,7 % der als ,,kernlos" klassifizierten Kreuzungsnachkommen konnten die Kernreste organoleptisch nicht wahrgenommen werden. K e y wo r d s: table grape, berry, seedlessness, stenospermocarpy, seed trace, crossing, genetics, segregation ratio, recessive gene. Introduction In numerous countries a definite consumer preference for seedless grapes is being evidenced. Because of the larger potential berry size, that can be further enhanced by gibberellin applications (WEAVER 1958), stenospermocarpic types are favored as table grapes to those showing stimulative parthenocarpy. In stenospermocarpic cultivars, pollination and fertilization are followed by var-ious degrees of embryo abortion (PEARSON 1932). Such cultivars have viable pollen and are used as male parents with normal seeded genotypes in order to produce some prog-eny with aborted seed. Analyzing the proportion of seeded to seedless progeny in such crosses, contradictory conclusions have been drawn (STOUT 1936, 1939; WEINBERGER and HARMON 1964; ÜLMO and BARIS 1973). lt has been also stated that back crosses to seed-less genotypes were not always effective. Crosses between seeded cultivars or selec-tions gave only 5 seedless hybrids out of a population of 1762 individuals (WEINBERGER and HARMON 1964). Hence the conclusion that seedlessness appears tobe a recessive character. We have carried out analyses of progenies from crosses between seeded and seedless parents during 1983, 1984, 1985, 1986 and 1988. Results and conclusions drawn are communicated in the present paper. Materials and methods Fruiting progeny from crosses between seeded parents and seedless parents used as male donors were analyzed and classified organoleptically by a panel of 3 judges into seeded (N) and seedless (SS). Due to variation in seed vestiges in seedless grapes, 80 P. SPIEGEL-ROY; Y. BARONand N. SAHAR the latter were divided into two groups: Small vestiges barely discernible in the pulp (S), and aborted seed with appreciable portions of sclerenchyma in the integuments, with the texture of the seed differing markedly from the texture of the pulp (B). Var-ious seedless male parents were used; the largest number of crosses were made using the following: Sultanina, Perlette, Flame Seedless and L12 (a large-berried type). The number of different seeded parents used was 13 in 1983, 17 in 1984, 13 in 1985, 13 in 1986 and 19 in 1988. The number of populations analyzed each year can be deduced from Table 1 (degrees of freedom for determination of homogeneity chi-square plus one). Statistical analysis included testing of hypothesis by the chi-square method, test for indepen-dence and calculation of homogeneity chi-square. Results The number of hybrids, seeded (N) and seedless (B + S) is given in Table 1. Assuming a theoretical distribution of 3 seeded: 1 seedless, x,2 values (after using Yates correction) are given for each year. In all cases x,2 values are small enough (probabili-ties > 0.05) and point to acceptance of hypothesis. During 4 years out of 5, homogeneity x,2 has been outside the accepted limit, signi-fying that the samples have not been homogeneous. Only in 1986 with a x,2 of 27 .77 and di. of 26, probability of the x,2 value was between 0.30 and 0.50. As each progeny has been analyzed separately for x,2, number of population with a x,2 that fits the assump-tion of a 3 : 1 ratio is also given. In all cases number of populations in which x,2 values have a probability > 0.05 clearly exceeds the number of populations in which the hypothesis of the 3 : 1 segregation had to be rejected. Table 1 Segregation of Vitis vinifera progeny for seedlessness from crosses between seeded (female) and stenospermocarpic (male) parents · Postulated segregation ratio 3 seeded: 1 seedless Die Aufspaltung des Merkmals ,,Kernlosigkeit" bei den Kreuzungsnachkommen aus kernhaltigen (weiblichen) und stenospermokarpen (männlichen) Vitis-vinifera-Sorten · Gefordertes Aufspal-tungsverhältnis „kernhaltig : kernlos" - 3 : 1 Seedless Homogeneity No. Normal Year indivi- seeded Noticeable Slight xz x2 d.f. xz > o.o5 X2 < 0.05 duals (N) seed traces seed (B) traces (S) accepted rejected 1983 483 364 85 34 0.03 68.86 23 20 4 1984 705 580 125 63 0.11 59.34 23 18 6 1985 661 545 116 49 1.08 76.20 24 17 8 1986 1240 1003 237 128 1.97 27.77 26 26 1 1988 1988 1457 391 140 3.01 80.50 22 16 7 We have also performed tests for independence (contingency x,2) comparing data of 1984 and 1985, and in another set those of 1986 and 1988 (Table 2). In both cases the calculated x,2 is significantly lower than the x! value at the 5 % level. Thus no interac-tion effect to cause differences in results is evident. While hybrid seedlings bearing Inheritance of seedlessness 81 fruit with seed traces have been pooled together with those yielding no noticeable seed traces in testing genetic ratlos of seeded vs seedless, their separate determination as Table 2 Contingency chi-square for data on segregation of progeny from seeded x seedless crosses of Vitis vinifera for seedless Kontingenz des x2-Tests für die Aufspaltung des Merkmals „Kernlosigkeit" von Kreuzungskombi-nationen ,,kernhaltig x kernlos" (Vitis vinifera) Year Seeded progeny Seedless progeny Contingency x2 1984 580 188 1.24 1985 545 165 1986 1003 365 0.04 1988 1457 531 performed is still of obvious practical significance. 'S' types (seedless) are clearly pre-ferred to 'B' (seedless types with noticeable seed traces). Table 3 summarizes the pro-portion of the two groups, indicating a preponderance of the group with seed traces. Thus, while the ratio of hybrids bearing seedless fruit from progenies of a seeded x seedless approximates 1 : 3 (Table 1), only about 30 % of the seedless progeny bore, under conditions prevailing in our breeding block, fruit without noticeable seed traces. This amounts to about 7.5 % only of the total progeny obtained (Table 3). Table 3 Proportion of progeny with seed traces barely noticeable (S) in total progeny rated as seedless (B + S) from crosses between seeded x seedless (Vitis vinifera) Anteil der Nachkommen mit kaum wahrnehmbaren Kernresten (S) an der gesamten als „kernlos" (B + S) klassifizierten Nachkommenschaft aus Kreuzungen „kernhaltig x kernlos" (Vitis vinifera) Numberof Progeny with barely noticeable seed traces Year seedless progeny Number Percentage 1983 119 34 28.6 1984 188 63 33.5 1985 165 49 29.7 1986 365 128 35.1 1988 531 140 26.4 A segregation of 3 : 1 between seeded and seedless in the total number of indivi-duals produced from seeded x seedless crosses of Vitis vinifera, each year, points to the 'seedless trait' as depending on two complementary recessive genes (aass). Proba-bly most, if not all seeded cultivars can thus be regarded as heterozygous dominants for the seeded condition (AaSs). This would yield in F1 from crosses between seeded and seedless a 3 : 1 ratio. 82 P. SPIEGEL-ROY; Y. BARON and N. SAHAR Discussion Crosses between seeded V. vinifera as a female parent and seedless types as a male parent have been known to yield a varying number of seedless progeny. Our work clearly shows that seedlessness is a recessive trait. Moreover, counts made during 5 nearly consecutive years (similar work in 1987 was not completed) show that a 3 : 1 ratio is being obtained, if results of all progenies derived from such a cross are pooled. Number of hybrids analyzed ranged from 483 to 1988. On the basis of the results obtained, inheritance of seedlessness seems to be determined by two complementary recessive genes. Nearly all seeded cultivars used as a parent yielded also seedless prog-eny and can thus be considered heterozygotes. However, some of the populations from certain crosses deviated sharply from the 3 : 1 ratio, causing high x2 values in 4 out of the 5 years examined. Examination of the performance of seedless parent used in the crosses (results not given) does not seem to reveal significant differences as to proportion of seedless prog-eny obtained, though differences in the proportion of seedless progeny with very slight seed traces to those with noticeable seed traces seem indicated. In contrast, crosses involving certain seeded parents do show differences in the proportion of seeded to seedless in the progeny. For an exact comparison tobe valid, a diallele setup and ana-lysis are required. If inheritance does depend on two complementary recessive genes, backcrosses to about 2/3 of the seeded parent derived from a seeded x seedless cross could show a more favorable segregation, increasing chances of seedlessness up to 50 %. Crosses of AaSs (seeded) and aass (seedless) would yield AaSs, aaSs, Aass and aass. Crosses ofAass or aaSs (though their phenotype will be indistinguishable from AaSa) would yield a 1 : 1 segregation instead of the 3 : 1 ratio of seeded : seedless when AaSa is used as a female parent. Similar crosses made in the past could have given a 1 : 1 ratio, originating the assumption that seedlessness behaves as a dominant trait (STOUT 1939; ÜLMO and BARIS 1973). All our crosses have been performed with stenospermocarpic male parents. Seed traces of varying size are obtained in the seedless progeny, ranging from nearly unno-ticeable traces smaller even or comparable to those obtained in Sultanina to large seed traces unaffected by gibberellin treatments during bloom. Hybrids with stenospermo-carpic fruit have been divided by us into two subclasses, based on extent and nature of seed traces in the fruit. The percentage of hybr~ds bearing fruit with small vestiges barely discernible in the pulp has been found to vary different years between 26.4 % and 35.1 % of the totality of the hybrids considered seedless. This would amount to only 6.5-8.3 % of the total progeny from seeded x seedless crosses. With the aid of the new in vitro method of culturing ovules and embryos of seedless parents (EMERSHAD and RAMMING 1984; SPIEGEL-ROY et al. 1985), a very high proportion of seedless progeny is being obtained from hybrids of crosses performed between two seedless parents. Genetic ratlos from such crosses will be reported in another paper. Summary Progenies derived from seeded (female parent) and seedless (male parent) Vitis vinifera have been analyzed during 5 years for number of hybrids bearing seeded vs seedless fruit. Distribution closely approximated a 3 : 1 ratio. x2 for the different years (1983, 1984, 1985, 1986, 1988) were 0.03, 0.11, 1.17, 1.97 and 3.01. Homogeneity of x2 was Inheritance of seedlessness 83 within the accepted Umit (0.3-0.5) only in 1986. Most of the populations during all years had xz conforming to the 3 : 1 ratio. Two tests for. independence during consecu-tive years showed no interaction. On the basis of these results, inheritance of seedless-ness is postulated to depend on two complementary recessive genes. Only 30.7 % of the progeny rated as seedless bore fruit without noticeable seed traces. Literature eited EMJ<:RSHAD, R. L.; RAMMING, D. W.; 1984: In ovulo embryo eulture of Vitis vinifera L. cv. Thompson Seedless. Amer. J. Bot. 71, 873-877. ÜLMO, H. P.; BAR!s, C.; 1973: Obtention des raisins de table apyrenes. Symp. Intern. Raisins de Table, Llmasol, juillet 1973; Doeument 20. PEARsoN, H. M.; 1932: Parthenocarpy and seed abortion in Vitis vinifera. Proc. Amer. Soc. Hort. Sei. 29, 169-175. SPIEGEL-RoY, P.; SAHAR, N.; BARON, I.; LAv1, U.; 1985: In vitro culture and plant formation from grape . cultivars with abortive ovules and seeds. J. Amer. Soc. Hort. Sei. 110, 109-112. Srour, A. B.; 1936: Seedlessness in grapes. New York Agricult. Exp. Sta. Tech. Bull. 238. ; 1939: Progress in breeding for seedless grapes. Proc. Amer. Soc. Hqrt. Sei. 37, 626-629. WEAVER, R.J.; 1958: Effect of gibberellie aeid on fruit set and berry enlargement in seedless grapes of Vitis vinifera. Nature 181, 851-852. WEINBERGER, J. H.; HARMON, F. N.; 1964: Seedlessness in vinifera grapes. Proe. Amer. Soc. Hort. Sei. 85, 270-274. Eingegangen am 17. 8. 1989 P. SPIEGEL-ROY Dept. of Fruit Breeding and Genetics Volcani Center, A.R.O. BetDagan Israel 

Inheritance of seedlessness in seeded x seedless progeny of Vitis vinifera L.

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Inheritance of seedlessness in seeded x seedless progeny of Vitis vinifera L.

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